刺梨的适钙类型及对高钙生境的适应性

doi:10.17521/cjpe.2022.0172

摘要/Abstract

摘要：

刺梨(Rosa roxburghii)是原产中国西南地区的特色经济树种, 贵州将其作为喀斯特山区重点发展的果树。迄今, 刺梨能否适应喀斯特地区高钙生境尚不清楚。明确刺梨的适钙类型及其对高钙生境的适应性, 旨在为刺梨种植区域的确定提供科学依据。该研究对贵州喀斯特地区不同钙生境下50个野生刺梨植株的不同器官和立地土壤进行取样, 测定土壤的pH、交换性钙、镁含量和植株不同器官中总钙、草酸钙及叶中的钙和镁的含量, 明确不同器官中草酸钙含量占总钙含量的比例, 分析土壤交换性钙、镁含量与不同器官总钙、叶片钙和镁含量的相关性关系, 观察低钙和高钙生境下不同器官中及叶表面草酸钙晶体的分布。对土壤的分析结果表明: 喀斯特地区野生刺梨立地土壤的pH高, 交换性钙、镁的含量高且差异大。在不同钙生境下, 野生刺梨不同器官中的总钙含量与土壤中交换性钙含量呈显著正相关关系, 叶片中总钙和镁的含量与土壤中交换性钙、镁含量的相关性也是如此。刺梨枝、叶是富集钙最多的器官, 根和果实中钙的含量相对较少, 所有样本叶片中总钙和镁的含量在1.71%-2.73%之间, 枝和叶中草酸钙含量占总钙含量的比例分别为55.81%和52.76%, 而根和果实中只占29.34%和34.30%。在高钙生境中, 野生刺梨枝、叶中和叶片气孔边缘有大量草酸钙棱晶, 果实中有少量针晶; 而在低钙生境中, 野生刺梨的根和果实中及叶片气孔边缘均无草酸钙晶体分布。该研究明确了刺梨的适钙类型属于中间型, 对喀斯特地区不同钙生境均有较强的适应性, 枝、叶中大量储存草酸钙和通过叶片气孔排钙的生理特性, 对降低刺梨体内钙离子浓度和缓解高钙胁迫对自身的伤害具有重要作用。研究结果对喀斯特地区刺梨种植区域的确定具有重要指导意义。

关键词: 刺梨, 喀斯特地区, 高钙生境, 适钙类型, 适应性分析

Abstract:

Aims Rosa roxburghii is a special economic tree species native to southwest China, and it has been taken as a key fruit tree in karst mountainous areas of Guizhou Province. However, it is unclear whether R. roxburghii can adapt to high-calcium habitats in the karst areas. To provide scientific evidence for determining the potential planting sites of R. roxburghii, the calcium-tolerance type and adaptability to high-calcium habitats of R. roxburghii were needed to be clarified.

Methods In this study, different organs from 50 R. roxburghii individuals and different calcareous soils where R. roxburghii grew were sampled in karst areas of Guizhou. The pH, exchangeable calcium and magnesium contents in the soil, total calcium and calcium oxalate contents in the plant organs, and calcium and magnesium contents in the leaves were determined, and the proportion of calcium oxalate in different organs to total calcium content was calculated. Then, the correlations between the exchangeable calcium and magnesium content in the soil with total calcium in different organs, and calcium and magnesium content in the leaves were analyzed. In addition, the distribution of calcium oxalate crystals in different organs as well as on the leaf surface was observed in samples collected from low-calcium and high-calcium habitats.

Important findings The soil at R. roxburghii sites in karst areas had a high pH, and the content of exchangeable calcium and magnesium was abundant and varies greatly. In different calcareous habitats, the total calcium content in different organs of R. roxburghii was significantly and positively correlated with the exchangeable calcium content in the soil. Similarly, the contents of total calcium and magnesium in the leaves were correlated with those in the soil. The branches and leaves were the organs that accumulated the most calcium, whereas the calcium enrichment in the roots and fruit were relatively low. The total calcium and magnesium contents of all the sample leaves ranged from 1.71% to 2.73%, among which, the proportions of calcium oxalate in branches and leaves were 55.81% and 52.76% of the total calcium, respectively; while only 29.34% and 34.30% were in roots and fruit. In samples from a high-calcium habitats, calcium oxalate prismatic crystals were abundant in the branches, leaves, and around stomatal pores, while few needle crystals were observed in the fruits. However, in samples from a low-calcium environment, no calcium oxalate crystals were observed in the roots, fruit, and leaf stomatal pores. In this study, we clarified that the R. roxburghii is an intermediate type of calcium-tolerance, which has strong adaptability to different calcium habitats in karst areas. The physiological characteristics of storing abundant calcium oxalate crystals in branches and leaves, and excreting calcium through leaf stomata play important roles in reducing the calcium concentration in R. roxburghii tissues and alleviating damages from high-calcium stress. The results of this study provide important guidance for the determination of suitable planting sites for R. roxburghii in karst areas.

Key words: Rosa roxburghii, karst area, high-calcium habitat, calcium-tolerance type, adaptability analysis

孟庆静, 樊卫国. 刺梨的适钙类型及对高钙生境的适应性. 植物生态学报, 2022, 46(12): 1562-1572. DOI: 10.17521/cjpe.2022.0172

MENG Qing-Jing, FAN Wei-Guo. Calcium-tolerance type and adaptability to high-calcium habitats of Rosa roxburghii. Chinese Journal of Plant Ecology, 2022, 46(12): 1562-1572. DOI: 10.17521/cjpe.2022.0172

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0172

https://www.plant-ecology.com/CN/Y2022/V46/I12/1562

图/表 9

表1 喀斯特地区野生刺梨立地土壤的pH和交换性钙、镁的含量(n = 50)

Table 1 Characteristics of pH, exchangeable calcium and magnesium content of wild Rosa roxburghii site soil in karst area (n = 50)

指标 Index	分级 Classification	样本数 Number (n)	范围 Range	占比 Percentage (%)	平均值 Mean	变异系数 Coefficient of variation (%)
pH	酸性 Acid (<5.5)	5	4.73-8.58	10.00	7.28	14.77
	微酸性 Faintly acid (5.5-6.5)	7		14.00
	中性 Neutral (6.5-7.5)	8		16.00
	碱性 Alkaline (7.5-8.5)	29		48.00
	强碱性 Highly alkaline (>8.5)	1		2.00
交换性钙含量 Exchangeable calcium content (mg·kg^-1)	缺乏 Lack (<250)	0	406.41-9 425.98	0.00	3 839.36	60.59
	适量 Suitable (250-1 000)	5		10.00
	高量 High (1 000-2 000)	5		10.00
	过量 Superfluous (>2 000)	40		80.00
交换性镁含量 Exchangeable magnesium content (mg·kg^-1)	缺乏 Lack (<50)	4	23.04-1 354.48	8.00	315.01	97.80
	适量 Suitable (50-100)	8		16.00
	高量 High (100-200)	13		26.00
	过量 Superfluous (>200)	25		50.00

表2 喀斯特地区野生刺梨根、枝、叶、果实中的总钙含量范围、变异系数及分布特征

Table 2 Range, coefficient of variation and distribution characteristics of total calcium content in root, branch, leaf and fruit of wild Rosa roxburghii in karst area

器官 Organ	范围 Range (g·kg^-1)	平均值 Mean (g·kg^-1)	标准差 Standard deviation	变异系数 Coefficient of variation (%)	总钙含量分布的样本数 Sample number of total calcium content distribution (n)
器官 Organ	范围 Range (g·kg^-1)	平均值 Mean (g·kg^-1)	标准差 Standard deviation	变异系数 Coefficient of variation (%)	<10 g·kg^-1	10-20 g·kg^-1	>20 g·kg^-1
根 Root	3.87-13.47	6.51^b	2.25	34.64	45	5	0
枝 Branch	11.01-24.96	19.92^a	3.27	16.41	0	25	25
叶 Leaf	13.95-24.95	19.72^a	2.96	15.01	0	28	22
果实 Fruit	3.21-11.50	6.19^b	1.81	29.22	27	2	0

图1 喀斯特地区50株野生刺梨叶片中钙和镁的含量(平均值±标准差)。

Fig. 1 Calcium and magnesium content in leaves of 50 wild Rosa roxburghii plants in karst area (mean ± SD).

图2 喀斯特地区野生刺梨叶片中钙和镁的含量与土壤中交换性钙和镁含量的相关性(n = 50)。**, p < 0.01。

Fig. 2 Correlation between calcium and magnesium content in leaf of wild Rosa roxburghii and exchangeable calcium and magnesium content in soil in karst area (n = 50). **, p < 0.01.

图3 喀斯特地区野生刺梨枝、叶、果实、根中的总钙含量与土壤中交换性钙含量的相关性。*, p < 0.05; **, p < 0.01。

Fig. 3 Correlation between total calcium content in branches, leaves, fruits, and roots of wild Rosa roxburghii and exchangeable calcium content in soil in karst area. *, p < 0.05; **, p < 0.01.

表3 喀斯特地区野生刺梨根、枝、叶、果实中的草酸钙含量范围、变异系数及其占各自总钙含量的百分比

Table 3 Content range, coefficient of variation and percentage of calcium oxalate in total calcium in root, branch, leaf and fruit of wild Rosa roxburghii in karst area

器官 Organ	草酸钙含量范围 Content range of calcium oxalate (g·kg^-1)	含量平均值 Content mean (g·kg^-1)	标准差 Standard deviation	变异系数 Coefficient of variation (%)	占器官总钙的百分比 Percentage of total calcium in organs (%)
根 Root	0.35-5.64	1.91^b	1.09	57.02	29.35^c
枝 Branch	3.34-16.88	11.12^a	2.82	25.39	55.81^a
叶 Leaf	7.16-13.67	10.41^a	1.64	15.73	52.76^a
果实 Fruit	0.52-4.96	2.12^b	1.01	47.52	34.30^b

表4 不同钙生境中土壤的交换性钙含量和野生刺梨植株不同器官中的总钙及草酸钙含量(平均值±标准差)

Table 4 Exchangeable calcium content in soil, total calcium and calcium oxalate content in different organs of wild Rosa roxburghii plants in different calcium conditions (mean ± SD)

生境 Condition	交换性钙含量 Exchangeable calcium content (mg·kg^-1)	总钙含量 Total calcium content (g·kg^-1)				草酸钙含量 Calcium oxalate content (g·kg^-1)
生境 Condition	交换性钙含量 Exchangeable calcium content (mg·kg^-1)	叶 Leaf	枝 Branch	果实 Fruit	根 Root	叶 Leaf	枝 Branch	果实 Fruit	根 Root
高钙 High calcium	5 938.12 ± 185.04^A	19.92 ± 0.70^A	20.90 ± 0.10^A	11.50 ± 0.14^A	10.07 ± 0.13^A	9.75 ± 0.14^A	10.10 ± 0.14^A	4.96 ± 0.02^A	2.34 ± 0.03^A
低钙 Low calcium	406.41 ± 14.41^B	14.07 ± 0.37^B	12.03 ± 0.08^B	3.86 ± 0.02^B	5.64 ± 0.13^B	7.53 ± 0.57^B	5.70 ± 0.23^B	0.52 ± 0.04^B	1.91 ± 0.06^B

图4 分别生长在两种钙生境中野生刺梨器官中及叶片气孔边缘的草酸钙晶体观察结果。A-F, 土壤交换性钙含量为5 938.12 mg·kg-1的高钙生境中野生刺梨不同器官内及叶面的草酸钙晶体观察结果。A, 叶片组织中的草酸钙棱晶。B, 枝韧皮部中的草酸钙棱晶。C, 果实组织中的草酸钙针晶。D, 根组织中没有草酸钙晶体。E, 气孔边缘的草酸钙堆积体。F, 气孔外的草酸钙棱晶。G-L, 土壤交换性钙含量为406.41 mg·kg-1的低钙生境中野生刺梨不同器官内及叶面的草酸钙晶体观察结果。G, 叶片组织中的草酸钙棱晶。H, 枝韧皮部中的草酸钙棱晶。I, 果实组织无草酸钙晶体。J, 根组织中无草酸钙晶体。K, 气孔周边无草酸钙堆积体。L, 在叶片表面及气孔周边无草酸钙晶体。

Fig. 4 Observation results of calcium oxalate crystals grown in the organs of Rosa roxburghii and at the stomata margins of leaves in two calcium conditions. A-F, Observation results of calcium oxalate crystals in different organs and leaves of R. roxburghii in high calcium conditions with soil exchangeable calcium content of 5 938.12 mg·kg-1. A, Calcium oxalate prism in leaf tissue. B, Calcium oxalate prism in branch phloem. C, Calcium oxalate raphid in fruit tissue. D, Root tissue without calcium oxalate crystals. E, Calcium oxalate deposits at the edge of the stomata. F, Calcium oxalate prism outside stomata. G-L, Observation results of calcium oxalate crystals in different organs and leaves of R. roxburghii in low calcium conditions with soil exchangeable calcium content of 406.41 mg·kg-1. G, Calcium oxalate prism in leaf tissue. H, Calcium oxalate prism in branch phloem. I, No calcium oxalate crystals were observed in fruit tissue. J, Root tissue without calcium oxalate crystals. K, No calcium oxalate deposits were observed around the stomata. L, No calcium oxalate crystals were observed around the stomata on the leaf surface.

图5 两种钙生境中的野生刺梨。左: 生长在pH 5.02、交换性钙含量为432 mg·kg-1的酸性黄壤中的野生刺梨, 植株周边生长有较多的酸性指示植物。右: 生长在pH 8.24、交换性钙含量为6 631 mg·kg-1的碱性石灰土中的野生刺梨。

Fig. 5 Rosa roxburghii in two calcium conditions. Left: R. roxburghii grows on acidic low calcium yellow soil with pH 5.02 and exchangeable calcium content of 432 mg·kg-1, there were more acidic indicator ferns around the plant. Right: R. roxburghii grows in alkaline calcareous soil with pH 8.24 and exchangeable calcium content up to 6 631 mg·kg-1.

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